Coronosaurus
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''Coronosaurus'' is a
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nom ...
of
centrosaurine Centrosaurinae (from the Greek, meaning "pointed lizards") is a subfamily of ceratopsid dinosaurs, a group of large quadrupedal ornithischians. Centrosaurine fossil remains are known primarily from the northern region of Laramidia (modern day A ...
ceratopsian Ceratopsia or Ceratopia ( or ; Ancient Greek, Greek: "horned faces") is a group of herbivore, herbivorous, beaked dinosaurs that thrived in what are now North America, Europe, and Asia, during the Cretaceous Period (geology), Period, although anc ...
dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the evolution of dinosaurs is t ...
s which lived in the
Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the younger of two epochs into which the Cretaceous Period is divided in the geologic time scale. Rock strata from this epoch form the Upper Cretaceous Series. The Cretaceous is named after ''creta'', ...
, in the middle
Campanian The Campanian is the fifth of six ages of the Late Cretaceous Epoch on the geologic timescale of the International Commission on Stratigraphy (ICS). In chronostratigraphy, it is the fifth of six stages in the Upper Cretaceous Series. Campani ...
stage. Its remains, two bone beds, were discovered by Phillip J. Currie in the Oldman Formation of
Alberta Alberta ( ) is one of the thirteen provinces and territories of Canada. It is part of Western Canada and is one of the three prairie provinces. Alberta is bordered by British Columbia to the west, Saskatchewan to the east, the Northwest T ...
,
Canada Canada is a country in North America. Its ten provinces and three territories extend from the Atlantic Ocean to the Pacific Ocean and northward into the Arctic Ocean, covering over , making it the world's second-largest country by to ...
, and its type and only species, ''Coronosaurus brinkmani'', was first described in 2005, as a new species within the genus '' Centrosaurus''. Later studies questioned the presence of a direct relationship, and in 2012 it was named as a separate genus. ''Coronosaurus'' means "crowned lizard", coming from "''corona''",
Latin Latin (, or , ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through ...
for crown, and "''sauros''",
Greek Greek may refer to: Greece Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group. *Greek language, a branch of the Indo-European language family. **Proto-Greek language, the assumed last common ancestor ...
for lizard; this name refers to the unique, crown-like shape of the horns on the top of its frill. Like other
ceratopsids Ceratopsidae (sometimes spelled Ceratopidae) is a family of ceratopsian dinosaurs including ''Triceratops'', ''Centrosaurus'', and ''Styracosaurus''. All known species were quadrupedal herbivores from the Upper Cretaceous. All but one species ar ...
, ''Coronosaurus'' had a large frill and horns on its head. These include a small pair of brow horns over its eyes, a large nasal horn on its snout, and, unique among ceratopsians, irregular, spiky bone masses on its frill. Growing up to around long and in weight, it was mid-sized for its kind. The genus is classified as a member of the Centrosaurini, a group of derived centrosaurines which has also been found include taxa such as ''
Styracosaurus ''Styracosaurus'' ( ; meaning "spiked lizard" from the Ancient Greek / "spike at the butt-end of a spear-shaft" and / "lizard") is a genus of herbivorous ceratopsian dinosaur from the Cretaceous Period (Campanian stage), about 75.5 to 74.5 ...
'', '' Spinops'', ''
Rubeosaurus ''Styracosaurus'' ( ; meaning "spiked lizard" from the Ancient Greek / "spike at the butt-end of a spear-shaft" and / "lizard") is a genus of herbivorous ceratopsian dinosaur from the Cretaceous Period (Campanian stage), about 75.5 to 74.5&nb ...
'', and '' Centrosaurus'', the genus it was originally placed within.


Discovery and naming

''Coronosaurus'' is known from two
bone bed A bone bed is any geological stratum or deposit that contains bones of whatever kind. Inevitably, such deposits are sedimentary in nature. Not a formal term, it tends to be used more to describe especially dense collections such as Lagerstätt ...
s, BB 138 and MRR BB, located in the upper unit of the Oldman Formation, of the Belly River Group, excavated by Philip Currie between 1996 and 2000.M.J. Ryan and A.P. Russell, 2003, "New centrosaurine ceratopsids from the late Campanian of Alberta and Montana and a review of contemporaneous and regional patterns of centrosaurine evolution", ''Journal of Vertebrate Paleontology'' 23(3): 91A Most of the ceratopsid material, if not all, from BB 138 in Dinosaur Provincial Park,
Alberta Alberta ( ) is one of the thirteen provinces and territories of Canada. It is part of Western Canada and is one of the three prairie provinces. Alberta is bordered by British Columbia to the west, Saskatchewan to the east, the Northwest T ...
, and the MRR BB near
Warner, Alberta Warner is a village in Alberta, Canada. It is surrounded by the County of Warner No. 5, approximately south of Lethbridge. Warner is a farming community. Warner is situated at the intersection of Highway 4 and Highway 36, about 38 km nor ...
, was referred to ''C. brinkmani''. Bone bed 138 is located approximately from
Brooks, Alberta Brooks is a city in southeast Alberta, Canada that is surrounded by the County of Newell. It is located on Highway 1 ( Trans-Canada Highway) and the Canadian Pacific Railway, approximately southeast of Calgary, and northwest of Medicine Hat. ...
, in the Oldman Formation and below the contact with the
Dinosaur Park Formation The Dinosaur Park Formation is the uppermost member of the Belly River Group (also known as the Judith River Group), a major geologic unit in southern Alberta. It was deposited during the Campanian stage of the Late Cretaceous, between about 76. ...
. The MRR BB located approximately southwest of BB 138, is also from the Oldman Formation. These bone beds date to the middle
Campanian The Campanian is the fifth of six ages of the Late Cretaceous Epoch on the geologic timescale of the International Commission on Stratigraphy (ICS). In chronostratigraphy, it is the fifth of six stages in the Upper Cretaceous Series. Campani ...
stage of the
Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the younger of two epochs into which the Cretaceous Period is divided in the geologic time scale. Rock strata from this epoch form the Upper Cretaceous Series. The Cretaceous is named after ''creta'', ...
period. Both the specimens and the precise localities are archived at the
Royal Tyrrell Museum of Palaeontology The Royal Tyrrell Museum of Palaeontology (RTMP, and often referred to as the Royal Tyrrell Museum) is a palaeontology museum and research facility in Drumheller, Alberta, Canada. The museum was named in honour of Joseph Burr Tyrrell, and is situ ...
, in Drumheller,
Alberta Alberta ( ) is one of the thirteen provinces and territories of Canada. It is part of Western Canada and is one of the three prairie provinces. Alberta is bordered by British Columbia to the west, Saskatchewan to the east, the Northwest T ...
. Michael J. Ryan and Anthony P. Russell described and named the
type species In zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the species that contains the biological type specim ...
, then as '' Centrosaurus brinkmani'', in
2005 File:2005 Events Collage V2.png, From top left, clockwise: Hurricane Katrina in the Gulf of Mexico; the Funeral of Pope John Paul II is held in Vatican City; " Me at the zoo", the first video ever to be uploaded to YouTube; Eris was discover ...
. Later studies, however, did not recover a monophyletic clade with the genus' type species ''Centrosaurus apertus'' in
phylogenetic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups ...
analyses. Due to this, Ryan,
David C. Evans David Cannon Evans (February 24, 1924 – October 3, 1998) was the founder of the computer science department at the University of Utah and co-founder (with Ivan Sutherland) of Evans & Sutherland, a pioneering firm in computer graphics hardwar ...
and Kieran M. Shepherd erected the genus ''Coronosaurus'' for the species in
2012 File:2012 Events Collage V3.png, From left, clockwise: The passenger cruise ship Costa Concordia lies capsized after the Costa Concordia disaster; Damage to Casino Pier in Seaside Heights, New Jersey as a result of Hurricane Sandy; People gat ...
. The generic name is derived from the
Latin Latin (, or , ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through ...
''corona'', meaning "crown" in reference to the multiple occurrences of extra epiparietals that cover the posterior margin of its parietal, giving it a crown-like appearance, and ''saurus'' (Latinized from
Greek Greek may refer to: Greece Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group. *Greek language, a branch of the Indo-European language family. **Proto-Greek language, the assumed last common ancestor ...
''sauros''), meaning "lizard". The specific name ''brinkmani'' honors Donald Brinkman, for his research in
palaeoecology Paleoecology (also spelled palaeoecology) is the study of interactions between organisms and/or interactions between organisms and their environments across geologic timescales. As a discipline, paleoecology interacts with, depends on and informs ...
of the Late Cretaceous environments of Alberta. In 2020, in response to the
COVID-19 pandemic The COVID-19 pandemic, also known as the coronavirus pandemic, is an ongoing global pandemic of coronavirus disease 2019 (COVID-19) caused by severe acute respiratory syndrome coronavirus 2 (SARS-CoV-2). The novel virus was first identi ...
, Ryan was asked about the naming of the dinosaur, and stated he came up with it because its frill ornamentation made him think of the corona of the sun. He also stated his fellow students at the time had jokingly called it ‘broccoli-ceratops’. The
holotype A holotype is a single physical example (or illustration) of an organism, known to have been used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of seve ...
of ''Coronosaurus'' is
TMP TMP can refer to any of the following: Chemistry * 2,2,6,6-Tetramethylpiperidine, an organic chemistry reagent * Thymidine monophosphate, a nucleotide * Trimethoprim, an antibiotic * Trimethyl phosphate, a solvent * Trimethylolpropane, a precurso ...
 2002.68.1. It is a large adult-sized parietal with an almost complete midline bar and a partial posterior bar with left P1–P3 processes and the partially eroded right P1-P2. The specimen lacks the extreme anterior margin of the midline bar that forms the posterior wall of the frontal fontanelle and the paper-thin lateral margins that define the medial margins of the supraorbital foramina. Other significant specimens according to Ryan & Russell (2005) include TMP 2002.68.3 (a parietal), TMP 2002.68.10 (a
postorbital The ''postorbital'' is one of the bones in vertebrate skulls which forms a portion of the dermal skull roof and, sometimes, a ring about the orbit. Generally, it is located behind the postfrontal and posteriorly to the orbital fenestra. In some ...
), and TMP 2002.68.5 ( supraorbitals).


Description

''Coronosaurus'' is a medium-sized
centrosaurine Centrosaurinae (from the Greek, meaning "pointed lizards") is a subfamily of ceratopsid dinosaurs, a group of large quadrupedal ornithischians. Centrosaurine fossil remains are known primarily from the northern region of Laramidia (modern day A ...
ceratopsid. Gregory S. Paul in 2010 estimated its body length at and its weight at .Paul, G.S., 2010, ''The Princeton Field Guide to Dinosaurs'', Princeton University Press p. 260 It had as an adult inflated supraorbital horncores — the "brow horns" above the eye sockets — but not elongated as in ''
Zuniceratops ''Zuniceratops'' ('Zuni-horned face') was a ceratopsian dinosaur from the mid Turonian of the Late Cretaceous Period of what is now New Mexico, United States. It lived about 10 million years earlier than the more familiar horned Ceratopsi ...
'', chasmosaurines, and more basal centrosaurines (like '' Albertaceratops'' and '' Diabloceratops''), that project laterally (to the sides) over the orbit. As a sub-adult its postorbital horncores are pyramidal in shape with a slight lateral inflection of the distal, upper, one half. Uniquely among ceratopsians, ''Coronosaurus'' possesses a number of accessory epiparietal
ossification Ossification (also called osteogenesis or bone mineralization) in bone remodeling is the process of laying down new bone material by cells named osteoblasts. It is synonymous with bone tissue formation. There are two processes resulting in ...
s rear parietal frill of the skull that fuse to the posterior and dorsal frill surface. They develop through
ontogeny Ontogeny (also ontogenesis) is the origination and development of an organism (both physical and psychological, e.g., moral development), usually from the time of fertilization of the egg to adult. The term can also be used to refer to the s ...
, the growth of the individual, as short spines that may fuse along their adjacent margins into larger, irregular bone masses. They are located close to the midline of the frill in a closely packed bunch above the base of the first epiparietal (P1), the bone spur that at each side growths downwards over the large opening in each frill half. They contribute to the substance of P1 and, through fusion, form a composite epiparietal, that is equivalent to the second epiparietal (P2) normally formed in a more lateral position. The P1 bases can thus be considered the growth positions of the second epiparietals, the P2 loci. ''Coronosaurus'' has also a uniquely shaped P3 epiparietal. It is variably developed as a short tongue-like hook or tapered spike that is oriented dorsolaterally, following the curve of the frill. Otherwise, as far as is known ''Coronosaurus'' resembles its relatives '' Centrosaurus'' and ''
Styracosaurus ''Styracosaurus'' ( ; meaning "spiked lizard" from the Ancient Greek / "spike at the butt-end of a spear-shaft" and / "lizard") is a genus of herbivorous ceratopsian dinosaur from the Cretaceous Period (Campanian stage), about 75.5 to 74.5 ...
'' in its morphology. For example, the
nasal Nasal is an adjective referring to the nose, part of human or animal anatomy. It may also be shorthand for the following uses in combination: * With reference to the human nose: ** Nasal administration, a method of pharmaceutical drug delivery * ...
of ''Coronosaurus'' closely resembles that of ''Centrosaurus apertus'' in its unfused juvenile or subadult and fused adult forms and appears to have undergone a similar ontogenetic changes. Its erect, laterally compressed nasal horn core has a blunt tip that is formed from the fusion of the opposing nasals. It sits over the posterior portion of the external nares, as it does in all centrosaurines. All juvenile and most adult specimens have gently recurved anterior and posterior margins resulting in most horns having an apex that is oriented at least slightly caudally (backwards).


Classification

In its original description, Ryan & Russell (2005) considered ''Coronosaurus'' to represent a new
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriat ...
of ''Centrosaurus'' on the basis of a small
phylogenetic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups ...
analysis. It included seventeen characters and nine taxa. ''Coronosaurus'' (as ''Centrosaurus brinkmani'') and ''C. apertus'' grouped at the base of the Centrosaurinae, while ''Styracosaurus'' was found to be more closely related to '' Pachyrhinosaurus'', and other derived centrosaurines, than to ''Centrosaurus''. In 2011, Anthony R. Fiorillo and Ronald S. Tykoski modified the analysis of Currie ''et al.'' (2008), with 54 characters, to include more taxa, such as ''C. brinkmani''. They found ''C. brinkmani'' to be the
sister taxon In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and ...
of the
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English ter ...
of ''Styracosaurus'' and ''C. apertus'', while '' Pachyrhinosaurus'' and other derived centrosaurines were in a separate lineage. Thus, this analysis found no support for the
monophyly In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic ...
of ''Centrosaurus''. Later, Farke ''et al.'' (2011) used 97 morphological characters to assess the phylogenetic position of a new
taxon In biology, a taxon ( back-formation from '' taxonomy''; plural taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular n ...
which they named '' Spinops''. ''Spinops'' was found to be in a
polytomy An internal node of a phylogenetic tree is described as a polytomy or multifurcation if (i) it is in a rooted tree and is linked to three or more child subtrees or (ii) it is in an unrooted tree and is attached to four or more branches. A tr ...
with ''Centrosaurus apertus'', ''C. brinkmani'' and ''Styracosaurus'', and therefore ''C. brinkmani'' was excluded from the analysis to increase the resolution. Finally, Ryan, Evans & Shepherd (2012) used the data matrix of Farke ''et al.'' (2011) to assess the phylogenetic position of ''
Xenoceratops ''Xenoceratops'' (meaning "alien horned face") is a genus of centrosaurine ceratopsid dinosaur known from the Late Cretaceous (middle Campanian stage), and is known to have lived in what is currently Alberta, Canada. The genus has one known speci ...
''. Their revised analysis had significantly better resolution than that presented by Farke ''et al.'' (2011), due in part to the additional scoring of missing characters for some taxa based on direct observation of their specimens. For example, five additional characters were scored for ''C. brinkmani''. In the resultant tree, it was found to be in more advanced position than ''Spinops'' and ''C. apertus'', as the sister taxon of ''Styracosaurus''. Thus the new generic name, ''Coronosaurus'', was given to it. The
cladogram A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to ...
presented below follows a phylogenetic analysis by Chiba ''et al.'' (2017):


Paleoecology

Studies suggest that the
paleoenvironment Paleoecology (also spelled palaeoecology) is the study of interactions between organisms and/or interactions between organisms and their environments across geologic timescales. As a discipline, paleoecology interacts with, depends on and informs ...
of the Oldman Formation was an ancient
coastal plain A coastal plain is flat, low-lying land adjacent to a sea coast. A fall line commonly marks the border between a coastal plain and a piedmont area. Some of the largest coastal plains are in Alaska and the southeastern United States. The Gulf Co ...
.Eberth, D.A., and Hamblin, A.P., 1993 , Tectonic, stratigraphic, and sedimentologic significance of a regional discontinuity in the upper Judith River Group (Belly River wedge) of southern Alberta, Saskatchewan, and northern Montana : Canadian Journal of Earth Sciences, 30, 174‒200. In addition to remains of ''Coronosaurus'', this formation has produced the remains of the theropods '' Saurornitholestes'', ''
Daspletosaurus ''Daspletosaurus'' ( ; meaning "frightful lizard") is a genus of tyrannosaurid dinosaur that lived in Laramidia between about 79.5 and 74 million years ago, during the Late Cretaceous Period. The genus ''Daspletosaurus'' contains three spec ...
'', '' Troodon'', '' Dromaeosaurus'' and ''
Hesperonychus ''Hesperonychus'' (meaning "western claw") was a small, carnivorous dinosaur. It was a member of the family Dromaeosauridae, along with its larger relatives ''Deinonychus'' and ''Velociraptor''. There is one described species, ''Hesperonychus eli ...
'', the ceratopsids '' Albertaceratops'', '' Chasmosaurus'', and ''
Anchiceratops ''Anchiceratops'' ( ) is an extinct genus of chasmosaurine ceratopsid dinosaur that lived approximately 72 to 71 million years ago during the latter part of the Cretaceous Period in what is now Alberta, Canada. ''Anchiceratops'' was a medium-siz ...
'', the hadrosaurids '' Brachylophosaurus'', '' Gryposaurus'', ''
Parasaurolophus ''Parasaurolophus'' (; meaning "near crested lizard" in reference to ''Saurolophus)'' is a genus of herbivorous hadrosaurid ornithopod dinosaur that lived in what is now North America and possibly Asia during the Late Cretaceous Period, abo ...
'', and ''
Corythosaurus ''Corythosaurus'' (; ) is a genus of hadrosaurid "duck-billed" dinosaur from the Upper Cretaceous Period, about 77–75.7 million years ago. It lived in what is now North America. Its name means "helmet lizard", derived from Greek κόρυ ...
'', the thescelosaurid '' Albertadromeus'' and the ankylosaurid '' Scolosaurus''.


See also

* Timeline of ceratopsian research


References

{{Taxonbar, from=Q135802 Fossils of Canada Centrosaurines Late Cretaceous dinosaurs of North America Fossil taxa described in 2012 Monotypic dinosaur genera Oldman fauna Paleontology in Alberta Campanian genus first appearances Campanian genus extinctions Ornithischian genera